The nerves attached to the brain are the cranial nerves, which are primarily responsible for the sensory and motor functions of the head and neck (one of these nerves targets organs in the thoracic and abdominal cavities as part of the parasympathetic nervous system). There are twelve cranial nerves, which are designated CNI through CNXII for “Cranial Nerve,” using Roman numerals for 1 through 12. They can be classified as sensory nerves, motor nerves, or a combination of both, meaning that the axons in these nerves originate out of sensory ganglia external to the cranium or motor nuclei within the brain stem. Sensory axons enter the brain to synapse in a nucleus. Motor axons connect to skeletal muscles of the head or neck. Three of the nerves are solely composed of sensory fibers; five are strictly motor; and the remaining four are mixed nerves.

Learning the cranial nerves is a tradition in anatomy courses, and students have always used mnemonic devices to remember the nerve names. A traditional mnemonic is the rhyming couplet, “On Old Olympus’ Towering Tops/A Finn And German Viewed Some Hops,” in which the initial letter of each word corresponds to the initial letter in the name of each nerve. The names of the nerves have changed over the years to reflect current usage and more accurate naming. An exercise to help learn this sort of information is to generate a mnemonic using words that have personal significance. The names of the cranial nerves are listed in image along with a brief description of their function, their source (sensory ganglion or motor nucleus), and their target (sensory nucleus or skeletal muscle). They are listed here with a brief explanation of each nerve (image).

The olfactory nerve and optic nerve are responsible for the sense of smell and vision, respectively. The oculomotor nerve is responsible for eye movements by controlling four of the extraocular muscles. It is also responsible for lifting the upper eyelid when the eyes point up, and for pupillary constriction. The trochlear nerve and the abducens nerve are both responsible for eye movement, but do so by controlling different extraocular muscles. The trigeminal nerve is responsible for cutaneous sensations of the face and controlling the muscles of mastication. The facial nerve is responsible for the muscles involved in facial expressions, as well as part of the sense of taste and the production of saliva. The vestibulocochlear nerve is responsible for the senses of hearing and balance. The glossopharyngeal nerve is responsible for controlling muscles in the oral cavity and upper throat, as well as part of the sense of taste and the production of saliva. The vagus nerve is responsible for contributing to homeostatic control of the organs of the thoracic and upper abdominal cavities. The spinal accessory nerve is responsible for controlling the muscles of the neck, along with cervical spinal nerves. The hypoglossal nerve is responsible for controlling the muscles of the lower throat and tongue.

Three of the cranial nerves also contain autonomic fibers, and a fourth is almost purely a component of the autonomic system. The oculomotor, facial, and glossopharyngeal nerves contain fibers that contact autonomic ganglia. The oculomotor fibers initiate pupillary constriction, whereas the facial and glossopharyngeal fibers both initiate salivation. The vagus nerve primarily targets autonomic ganglia in the thoracic and upper abdominal cavities.

Another important aspect of the cranial nerves that lends itself to a mnemonic is the functional role each nerve plays. The nerves fall into one of three basic groups. They are sensory, motor, or both (see image). The sentence, “Some Say Marry Money But My Brother Says Brains Beauty Matter More,” corresponds to the basic function of each nerve. The first, second, and eighth nerves are purely sensory: the olfactory (CNI), optic (CNII), and vestibulocochlear (CNVIII) nerves. The three eye-movement nerves are all motor: the oculomotor (CNIII), trochlear (CNIV), and abducens (CNVI). The spinal accessory (CNXI) and hypoglossal (CNXII) nerves are also strictly motor. The remainder of the nerves contain both sensory and motor fibers. They are the trigeminal (CNV), facial (CNVII), glossopharyngeal (CNIX), and vagus (CNX) nerves. The nerves that convey both are often related to each other. The trigeminal and facial nerves both concern the face; one concerns the sensations and the other concerns the muscle movements. The facial and glossopharyngeal nerves are both responsible for conveying gustatory, or taste, sensations as well as controlling salivary glands. The vagus nerve is involved in visceral responses to taste, namely the gag reflex. This is not an exhaustive list of what these combination nerves do, but there is a thread of relation between them.

The cranial nerves are composed of twelve pairs of nerves that emanate from the nervous tissue of the brain. In order to reach their targets they must ultimately exit/enter the cranium through openings in the skull. Hence, their name is derived from their association with the cranium. The following are the list of cranial nerves, their functions, and tumor examples:

Tastes and Odors

Both taste and odor stimuli are molecules taken in from the environment. The primary tastes detected by humans are sweet, sour, bitter, salty, and umami. The first four tastes need little explanation. The identification of umami as a fundamental taste occurred fairly recently—it was identified in 1908 by Japanese scientist Kikunae Ikeda while he worked with seaweed broth, but it was not widely accepted as a taste that could be physiologically distinguished until many years later. The taste of umami, also known as savoriness, is attributable to the taste of the amino acid L-glutamate. In fact, monosodium glutamate, or MSG, is often used in cooking to enhance the savory taste of certain foods. What is the adaptive value of being able to distinguish umami? Savory substances tend to be high in protein.

All odors that we perceive are molecules in the air we breathe. If a substance does not release molecules into the air from its surface, it has no smell. And if a human or other animal does not have a receptor that recognizes a specific molecule, then that molecule has no smell. Humans have about 350 olfactory receptor subtypes that work in various combinations to allow us to sense about 10,000 different odors. Compare that to mice, for example, which have about 1,300 olfactory receptor types, and therefore probably sense more odors. Both odors and tastes involve molecules that stimulate specific chemoreceptors. Although humans commonly distinguish taste as one sense and smell as another, they work together to create the perception of flavor. A person’s perception of flavor is reduced if he or she has congested nasal passages.

Reception and Transduction

Odorants (odor molecules) enter the nose and dissolve in the olfactory epithelium, the mucosa at the back of the nasal cavity (as illustrated in Figure). The olfactory epithelium is a collection of specialized olfactory receptors in the back of the nasal cavity that spans an area about 5 cm² in humans. Recall that sensory cells are neurons. An olfactory receptor, which is a dendrite of a specialized neuron, responds when it binds certain molecules inhaled from the environment by sending impulses directly to the olfactory bulb of the brain. Humans have about 12 million olfactory receptors, distributed among hundreds of different receptor types that respond to different odors. Twelve million seems like a large number of receptors, but compare that to other animals: rabbits have about 100 million, most dogs have about 1 billion, and bloodhounds—dogs selectively bred for their sense of smell—have about 4 billion. The overall size of the olfactory epithelium also differs between species, with that of bloodhounds, for example, being many times larger than that of humans.

Olfactory neurons are bipolar neurons (neurons with two processes from the cell body). Each neuron has a single dendrite buried in the olfactory epithelium, and extending from this dendrite are 5 to 20 receptor-laden, hair-like cilia that trap odorant molecules. The sensory receptors on the cilia are proteins, and it is the variations in their amino acid chains that make the receptors sensitive to different odorants. Each olfactory sensory neuron has only one type of receptor on its cilia, and the receptors are specialized to detect specific odorants, so the bipolar neurons themselves are specialized. When an odorant binds with a receptor that recognizes it, the sensory neuron associated with the receptor is stimulated. Olfactory stimulation is the only sensory information that directly reaches the cerebral cortex, whereas other sensations are relayed through the thalamus.

In the human olfactory system, (a) bipolar olfactory neurons extend from (b) the olfactory epithelium, where olfactory receptors are located, to the olfactory bulb. (credit: modification of work by Patrick J. Lynch, medical illustrator; C. Carl Jaffe, MD, cardiologist)

Smell and Taste in the Brain

Olfactory neurons project from the olfactory epithelium to the olfactory bulb as thin, unmyelinated axons. The olfactory bulb is composed of neural clusters called glomeruli, and each glomerulus receives signals from one type of olfactory receptor, so each glomerulus is specific to one odorant. From glomeruli, olfactory signals travel directly to the olfactory cortex and then to the frontal cortex and the thalamus. Recall that this is a different path from most other sensory information, which is sent directly to the thalamus before ending up in the cortex. Olfactory signals also travel directly to the amygdala, thereafter reaching the hypothalamus, thalamus, and frontal cortex. The last structure that olfactory signals directly travel to is a cortical center in the temporal lobe structure important in spatial, autobiographical, declarative, and episodic memories. Olfaction is finally processed by areas of the brain that deal with memory, emotions, reproduction, and thought.

Taste neurons project from taste cells in the tongue, esophagus, and palate to the medulla, in the brainstem. From the medulla, taste signals travel to the thalamus and then to the primary gustatory cortex. Information from different regions of the tongue is segregated in the medulla, thalamus, and cortex.

Olfaction (Smell)

Like taste, the sense of smell, or olfaction, is also responsive to chemical stimuli. The olfactory receptor neurons are located in a small region within the superior nasal cavity (Figure below). This region is referred to as the olfactory epithelium and contains bipolar sensory neurons. Each olfactory sensory neuron has dendrites that extend from the apical surface of the epithelium into the mucus lining the cavity. As airborne molecules are inhaled through the nose, they pass over the olfactory epithelial region and dissolve into the mucus. These odorant molecules bind to proteins that keep them dissolved in the mucus and help transport them to the olfactory dendrites. The odorant–protein complex binds to a receptor protein within the cell membrane of an olfactory dendrite. These receptors are G protein–coupled, and will produce a graded membrane potential in the olfactory neurons.

The axon of an olfactory neuron extends from the basal surface of the epithelium, through an olfactory foramen in the cribriform plate of the ethmoid bone, and into the brain. The group of axons called the olfactory tract connect to the olfactory bulb on the ventral surface of the frontal lobe. From there, the axons split to travel to several brain regions. Some travel to the cerebrum, specifically to the primary olfactory cortex that is located in the inferior and medial areas of the temporal lobe. Others project to structures within the limbic system and hypothalamus, where smells become associated with long-term memory and emotional responses. This is how certain smells trigger emotional memories, such as the smell of food associated with one’s birthplace. Smell is the one sensory modality that does not synapse in the thalamus before connecting to the cerebral cortex. This intimate connection between the olfactory system and the cerebral cortex is one reason why smell can be a potent trigger of memories and emotion.

The nasal epithelium, including the olfactory cells, can be harmed by airborne toxic chemicals. Therefore, the olfactory neurons are regularly replaced within the nasal epithelium, after which the axons of the new neurons must find their appropriate connections in the olfactory bulb. These new axons grow along the axons that are already in place in the cranial nerve.

The Olfactory System (a) The olfactory system begins in the peripheral structures of the nasal cavity. (b) The olfactory receptor neurons are within the olfactory epithelium. (c) Axons of the olfactory receptor neurons project through the cribriform plate of the ethmoid bone and synapse with the neurons of the olfactory bulb (tissue source: simian). LM × 812. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)

Higher Processing

The myelinated axons of ganglion cells make up the optic nerves. Within the nerves, different axons carry different qualities of the visual signal. Some axons constitute the magnocellular (big cell) pathway, which carries information about form, movement, depth, and differences in brightness. Other axons constitute the parvocellular (small cell) pathway, which carries information on color and fine detail. Some visual information projects directly back into the brain, while other information crosses to the opposite side of the brain. This crossing of optical pathways produces the distinctive optic chiasma (Greek, for “crossing”) found at the base of the brain and allows us to coordinate information from both eyes.

Once in the brain, visual information is processed in several places, and its routes reflect the complexity and importance of visual information to humans and other animals. One route takes the signals to the thalamus, which serves as the routing station for all incoming sensory impulses except olfaction. In the thalamus, the magnocellular and parvocellular distinctions remain intact, and there are different layers of the thalamus dedicated to each. When visual signals leave the thalamus, they travel to the primary visual cortex at the rear of the brain. From the visual cortex, the visual signals travel in two directions. One stream that projects to the parietal lobe, in the side of the brain, carries magnocellular (“where”) information. A second stream projects to the temporal lobe and carries both magnocellular (“where”) and parvocellular (“what”) information.

Another important visual route is a pathway from the retina to the superior colliculus in the midbrain, where eye movements are coordinated and integrated with auditory information. Finally, there is the pathway from the retina to the suprachiasmatic nucleus (SCN) of the hypothalamus. The SCN is a cluster of cells that is considered to be the body’s internal clock, which controls our circadian (day-long) cycle. The SCN sends information to the pineal gland, which is important in sleep/wake patterns and annual cycles.

ANATOMY OF THE VISUAL SYSTEM

The anatomy of the eye is illustrated in this diagram.

After passing through the pupil, light crosses the lens, a curved, transparent structure that serves to provide additional focus. The lens is attached to muscles that can change its shape to aid in focusing light that is reflected from near or far objects. In a normal-sighted individual, the lens will focus images perfectly on a small indentation in the back of the eye known as the fovea, which is part of the retina, the light-sensitive lining of the eye. The fovea contains densely packed specialized photoreceptor cells (Figure). These photoreceptor cells, known as cones, are light-detecting cells. The cones are specialized types of photoreceptors that work best in bright light conditions. Cones are very sensitive to acute detail and provide tremendous spatial resolution. They also are directly involved in our ability to perceive color.

While cones are concentrated in the fovea, where images tend to be focused, rods, another type of photoreceptor, are located throughout the remainder of the retina. Rods are specialized photoreceptors that work well in low light conditions, and while they lack the spatial resolution and color function of the cones, they are involved in our vision in dimly lit environments as well as in our perception of movement on the periphery of our visual field.

The two types of photoreceptors are shown in this image. Rods are colored green and cones are blue.

We have all experienced the different sensitivities of rods and cones when making the transition from a brightly lit environment to a dimly lit environment. Imagine going to see a blockbuster movie on a clear summer day. As you walk from the brightly lit lobby into the dark theater, you notice that you immediately have difficulty seeing much of anything. After a few minutes, you begin to adjust to the darkness and can see the interior of the theater. In the bright environment, your vision was dominated primarily by cone activity. As you move to the dark environment, rod activity dominates, but there is a delay in transitioning between the phases. If your rods do not transform light into nerve impulses as easily and efficiently as they should, you will have difficulty seeing in dim light, a condition known as night blindness.

Rods and cones are connected (via several interneurons) to retinal ganglion cells. Axons from the retinal ganglion cells converge and exit through the back of the eye to form the optic nerve. The optic nerve carries visual information from the retina to the brain. There is a point in the visual field called the blind spot: Even when light from a small object is focused on the blind spot, we do not see it. We are not consciously aware of our blind spots for two reasons: First, each eye gets a slightly different view of the visual field; therefore, the blind spots do not overlap. Second, our visual system fills in the blind spot so that although we cannot respond to visual information that occurs in that portion of the visual field, we are also not aware that information is missing.

The optic nerve from each eye merges just below the brain at a point called the optic chiasm. As Figure shows, the optic chiasm is an X-shaped structure that sits just below the cerebral cortex at the front of the brain. At the point of the optic chiasm, information from the right visual field (which comes from both eyes) is sent to the left side of the brain, and information from the left visual field is sent to the right side of the brain.

This illustration shows the optic chiasm at the front of the brain and the pathways to the occipital lobe at the back of the brain, where visual sensations are processed into meaningful perceptions.

Once inside the brain, visual information is sent via a number of structures to the occipital lobe at the back of the brain for processing. Visual information might be processed in parallel pathways which can generally be described as the “what pathway” and the “where/how” pathway. The “what pathway” is involved in object recognition and identification, while the “where/how pathway” is involved with location in space and how one might interact with a particular visual stimulus (Milner & Goodale, 2008; Ungerleider & Haxby, 1994). For example, when you see a ball rolling down the street, the “what pathway” identifies what the object is, and the “where/how pathway” identifies its location or movement in space.

The oculomotor nerve is responsible for eye movements by controlling four of the extraocular muscles. It is also responsible for lifting the upper eyelid when the eyes point up, and for pupillary constriction.

The trochlear nerve controls the superior oblique muscle to rotate the eye along its axis in the orbit medially, which is called intorsion, and is a component of focusing the eyes on an object close to the face.

The 5th and largest cranial nerve. Thetrigeminal nerveis a mixed motor and sensory nerve. The larger sensory part forms the ophthalmic, mandibular, and maxillary nerves which carry afferents sensitive to external or internal stimuli from the skin, muscles, and joints of the face and mouth and from the teeth. Most of these fibers originate from cells of the TRIGEMINAL GANGLION and project to the TRIGEMINAL NUCLEUS of the brain stem. The smaller motor part arises from the brain stem trigeminal motor nucleus and innervates the muscles of mastication.

The 6th cranial nerve which originates in the ABDUCENS NUCLEUS of the PONS and sends motor fibers to the lateral rectus muscles of the EYE. Damage to the nerve or its nucleus disrupts horizontal eye movement control.

Nerves of the Face and Oral Cavity

An iconic part of a doctor’s visit is the inspection of the oral cavity and pharynx, suggested by the directive to “open your mouth and say ‘ah.’” This is followed by inspection, with the aid of a tongue depressor, of the back of the mouth, or the opening of the oral cavity into the pharynx known as the fauces. Whereas this portion of a medical exam inspects for signs of infection, such as in tonsillitis, it is also the means to test the functions of the cranial nerves that are associated with the oral cavity.

The facial and glossopharyngeal nerves convey gustatory stimulation to the brain. Testing this is as simple as introducing salty, sour, bitter, or sweet stimuli to either side of the tongue. The patient should respond to the taste stimulus before retracting the tongue into the mouth. Stimuli applied to specific locations on the tongue will dissolve into the saliva and may stimulate taste buds connected to either the left or right of the nerves, masking any lateral deficits. Along with taste, the glossopharyngeal nerve relays general sensations from the pharyngeal walls. These sensations, along with certain taste stimuli, can stimulate the gag reflex. If the examiner moves the tongue depressor to contact the lateral wall of the fauces, this should elicit the gag reflex. Stimulation of either side of the fauces should elicit an equivalent response. The motor response, through contraction of the muscles of the pharynx, is mediated through the vagus nerve. Normally, the vagus nerve is considered autonomic in nature. The vagus nerve directly stimulates the contraction of skeletal muscles in the pharynx and larynx to contribute to the swallowing and speech functions. Further testing of vagus motor function has the patient repeating consonant sounds that require movement of the muscles around the fauces. The patient is asked to say “lah-kah-pah” or a similar set of alternating sounds while the examiner observes the movements of the soft palate and arches between the palate and tongue.

The facial and glossopharyngeal nerves are also responsible for the initiation of salivation. Neurons in the salivary nuclei of the medulla project through these two nerves as preganglionic fibers, and synapse in ganglia located in the head. The parasympathetic fibers of the facial nerve synapse in the pterygopalatine ganglion, which projects to the submandibular gland and sublingual gland. The parasympathetic fibers of the glossopharyngeal nerve synapse in the otic ganglion, which projects to the parotid gland. Salivation in response to food in the oral cavity is based on a visceral reflex arc within the facial or glossopharyngeal nerves. Other stimuli that stimulate salivation are coordinated through the hypothalamus, such as the smell and sight of food.

The 8th cranial nerve. The vestibulocochlear nerve has a cochlear part (COCHLEAR NERVE) which is concerned with hearing and a vestibular part (VESTIBULAR NERVE) which mediates the sense of balance and head position. The fibers of the cochlear nerve originate from neurons of the SPIRAL GANGLION and project to the cochlear nuclei (COCHLEAR NUCLEUS). The fibers of the vestibular nerve arise from neurons of Scarpa's ganglion and project to the VESTIBULAR NUCLEI.

Audition (Hearing)

Hearing, or audition, is the transduction of sound waves into a neural signal that is made possible by the structures of the ear (Figure below). The large, fleshy structure on the lateral aspect of the head is known as the auricle. Some sources will also refer to this structure as the pinna, though that term is more appropriate for a structure that can be moved, such as the external ear of a cat. The C-shaped curves of the auricle direct sound waves toward the auditory canal. The canal enters the skull through the external auditory meatus of the temporal bone. At the end of the auditory canal is the tympanic membrane, or ear drum, which vibrates after it is struck by sound waves. The auricle, ear canal, and tympanic membrane are often referred to as the external ear. The middle ear consists of a space spanned by three small bones called the ossicles. The three ossicles are the malleus, incus, and stapes, which are Latin names that roughly translate to hammer, anvil, and stirrup. The malleus is attached to the tympanic membrane and articulates with the incus. The incus, in turn, articulates with the stapes. The stapes is then attached to the inner ear, where the sound waves will be transduced into a neural signal. The middle ear is connected to the pharynx through the Eustachian tube, which helps equilibrate air pressure across the tympanic membrane. The tube is normally closed but will pop open when the muscles of the pharynx contract during swallowing or yawning.

Structures of the Ear. The external ear contains the auricle, ear canal, and tympanic membrane. The middle ear contains the ossicles and is connected to the pharynx by the Eustachian tube. The inner ear contains the cochlea and vestibule, which are responsible for audition and equilibrium, respectively.

The inner ear is often described as a bony labyrinth, as it is composed of a series of canals embedded within the temporal bone. It has two separate regions, the cochlea and the vestibule, which are responsible for hearing and balance, respectively. The neural signals from these two regions are relayed to the brain stem through separate fiber bundles. However, these two distinct bundles travel together from the inner ear to the brain stem as the vestibulocochlear nerve. Sound is transduced into neural signals within the cochlear region of the inner ear, which contains the sensory neurons of the spiral ganglia. These ganglia are located within the spiral-shaped cochlea of the inner ear. The cochlea is attached to the stapes through the oval window.

The oval window is located at the beginning of a fluid-filled tube within the cochlea called the scala vestibuli. The scala vestibuli extends from the oval window, travelling above the cochlear duct, which is the central cavity of the cochlea that contains the sound-transducing neurons. At the uppermost tip of the cochlea, the scala vestibuli curves over the top of the cochlear duct. The fluid-filled tube, now called the scala tympani, returns to the base of the cochlea, this time travelling under the cochlear duct. The scala tympani ends at the round window, which is covered by a membrane that contains the fluid within the scala. As vibrations of the ossicles travel through the oval window, the fluid of the scala vestibuli and scala tympani moves in a wave-like motion. The frequency of the fluid waves match the frequencies of the sound waves (Figure below). The membrane covering the round window will bulge out or pucker in with the movement of the fluid within the scala tympani.

Transmission of Sound Waves to Cochlea. A sound wave causes the tympanic membrane to vibrate. This vibration is amplified as it moves across the malleus, incus, and stapes. The amplified vibration is picked up by the oval window causing pressure waves in the fluid of the scala vestibuli and scala tympani. The complexity of the pressure waves is determined by the changes in amplitude and frequency of the sound waves entering the ear.

A cross-sectional view of the cochlea shows that the scala vestibuli and scala tympani run along both sides of the cochlear duct (Figure below). The cochlear duct contains several organs of Corti, which tranduce the wave motion of the two scala into neural signals. The organs of Corti lie on top of the basilar membrane, which is the side of the cochlear duct located between the organs of Corti and the scala tympani. As the fluid waves move through the scala vestibuli and scala tympani, the basilar membrane moves at a specific spot, depending on the frequency of the waves. Higher frequency waves move the region of the basilar membrane that is close to the base of the cochlea. Lower frequency waves move the region of the basilar membrane that is near the tip of the cochlea.

Cross Section of the Cochlea. The three major spaces within the cochlea are highlighted. The scala tympani and scala vestibuli lie on either side of the cochlear duct. The organ of Corti, containing the mechanoreceptor hair cells, is adjacent to the scala tympani, where it sits atop the basilar membrane.

The organs of Corti contain hair cells, which are named for the hair-like stereocilia extending from the cell’s apical surfaces (Figure below). The stereocilia are an array of microvilli-like structures arranged from tallest to shortest. Protein fibers tether adjacent hairs together within each array, such that the array will bend in response to movements of the basilar membrane. The stereocilia extend up from the hair cells to the overlying tectorial membrane, which is attached medially to the organ of Corti. When the pressure waves from the scala move the basilar membrane, the tectorial membrane slides across the stereocilia. This bends the stereocilia either toward or away from the tallest member of each array. When the stereocilia bend toward the tallest member of their array, tension in the protein tethers opens ion channels in the hair cell membrane. This will depolarize the hair cell membrane, triggering nerve impulses that travel down the afferent nerve fibers attached to the hair cells. When the stereocilia bend toward the shortest member of their array, the tension on the tethers slackens and the ion channels close. When no sound is present, and the stereocilia are standing straight, a small amount of tension still exists on the tethers, keeping the membrane potential of the hair cell slightly depolarized.

Hair Cell. The hair cell is a mechanoreceptor with an array of stereocilia emerging from its apical surface. The stereocilia are tethered together by proteins that open ion channels when the array is bent toward the tallest member of their array, and closed when the array is bent toward the shortest member of their array.

Cochlea and Organ of Corti LM × 412. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)

Equilibrium (Balance)

Along with audition, the inner ear is responsible for encoding information about equilibrium, the sense of balance. A similar mechanoreceptor—a hair cell with stereocilia—senses head position, head movement, and whether our bodies are in motion. These cells are located within the vestibule of the inner ear. Head position is sensed by the utricle and saccule, whereas head movement is sensed by the semicircular canals. The neural signals generated in the vestibular ganglion are transmitted through the vestibulocochlear nerve to the brain stem and cerebellum.

The utricle and saccule are both largely composed of macula tissue (plural = maculae). The macula is composed of hair cells surrounded by support cells. The stereocilia of the hair cells extend into a viscous gel called the otolithic membrane (Figure). On top of the otolithic membrane is a layer of calcium carbonate crystals, called otoliths. The otoliths essentially make the otolithic membrane top-heavy. The otolithic membrane moves separately from the macula in response to head movements. Tilting the head causes the otolithic membrane to slide over the macula in the direction of gravity. The moving otolithic membrane, in turn, bends the sterocilia, causing some hair cells to depolarize as others hyperpolarize. The exact position of the head is interpreted by the brain based on the pattern of hair-cell depolarization.

Linear Acceleration Coding by Maculae The maculae are specialized for sensing linear acceleration, such as when gravity acts on the tilting head, or if the head starts moving in a straight line. The difference in inertia between the hair cell stereocilia and the otolithic membrane in which they are embedded leads to a shearing force that causes the stereocilia to bend in the direction of that linear acceleration.

The semicircular canals are three ring-like extensions of the vestibule. One is oriented in the horizontal plane, whereas the other two are oriented in the vertical plane. The anterior and posterior vertical canals are oriented at approximately 45 degrees relative to the sagittal plane (Figure). The base of each semicircular canal, where it meets with the vestibule, connects to an enlarged region known as the ampulla. The ampulla contains the hair cells that respond to rotational movement, such as turning the head while saying “no.” The stereocilia of these hair cells extend into the cupula, a membrane that attaches to the top of the ampulla. As the head rotates in a plane parallel to the semicircular canal, the fluid lags, deflecting the cupula in the direction opposite to the head movement. The semicircular canals contain several ampullae, with some oriented horizontally and others oriented vertically. By comparing the relative movements of both the horizontal and vertical ampullae, the vestibular system can detect the direction of most head movements within three-dimensional (3-D) space.

Rotational Coding by Semicircular Canals Rotational movement of the head is encoded by the hair cells in the base of the semicircular canals. As one of the canals moves in an arc with the head, the internal fluid moves in the opposite direction, causing the cupula and stereocilia to bend. The movement of two canals within a plane results in information about the direction in which the head is moving, and activation of all six canals can give a very precise indication of head movement in three dimensions.

The 9th cranial nerve. The glossopharyngeal nerve is a mixed motor and sensory nerve; it conveys somatic and autonomic efferents as well as general, special, and visceral afferents. Among the connections are motor fibers to the stylopharyngeus muscle, parasympathetic fibers to the parotid glands, general and taste afferents from the posterior third of the tongue, the nasopharynx, and the palate, and afferents from baroreceptors and CHEMORECEPTOR CELLS of the carotid sinus.

Only a few recognized submodalities exist within the sense of taste, or gustation. Until recently, only four tastes were recognized: sweet, salty, sour, and bitter. Research at the turn of the 20th century led to recognition of the fifth taste, umami, during the mid-1980s. Umami is a Japanese word that means “delicious taste,” and is often translated to mean savory. Very recent research has suggested that there may also be a sixth taste for fats, or lipids.

Gustation is the special sense associated with the tongue. The surface of the tongue, along with the rest of the oral cavity, is lined by a stratified squamous epithelium. Raised bumps called papillae (singular = papilla) contain the structures for gustatory transduction. There are four types of papillae, based on their appearance (Figure): circumvallate, foliate, filiform, and fungiform. Within the structure of the papillae are taste buds that contain specialized gustatory receptor cells for the transduction of taste stimuli. These receptor cells are sensitive to the chemicals contained within foods that are ingested, and they release neurotransmitters based on the amount of the chemical in the food. Neurotransmitters from the gustatory cells can activate sensory neurons in the facial, glossopharyngeal, and vagus cranial nerves.

Salty taste is simply the perception of sodium ions (Na⁺) in the saliva. When you eat something salty, the salt crystals dissociate into the component ions Na⁺ and Cl^–, which dissolve into the saliva in your mouth. The Na⁺ concentration becomes high outside the gustatory cells, creating a strong concentration gradient that drives the diffusion of the ion into the cells. The entry of Na⁺ into these cells results in the depolarization of the cell membrane and the generation of a receptor potential.

Sour taste is the perception of H⁺ concentration. Just as with sodium ions in salty flavors, these hydrogen ions enter the cell and trigger depolarization. Sour flavors are, essentially, the perception of acids in our food. Increasing hydrogen ion concentrations in the saliva (lowering saliva pH) triggers progressively stronger graded potentials in the gustatory cells. For example, orange juice—which contains citric acid—will taste sour because it has a pH value of approximately 3. Of course, it is often sweetened so that the sour taste is masked.

The first two tastes (salty and sour) are triggered by the cations Na⁺ and H⁺. The other tastes result from food molecules binding to a G protein–coupled receptor. A G protein signal transduction system ultimately leads to depolarization of the gustatory cell. The sweet taste is the sensitivity of gustatory cells to the presence of glucose dissolved in the saliva. Other monosaccharides such as fructose, or artificial sweeteners such as aspartame (NutraSweet™), saccharine, or sucralose (Splenda™) also activate the sweet receptors. The affinity for each of these molecules varies, and some will taste sweeter than glucose because they bind to the G protein–coupled receptor differently.

Bitter taste is similar to sweet in that food molecules bind to G protein–coupled receptors. However, there are a number of different ways in which this can happen because there are a large diversity of bitter-tasting molecules. Some bitter molecules depolarize gustatory cells, whereas others hyperpolarize gustatory cells. Likewise, some bitter molecules increase G protein activation within the gustatory cells, whereas other bitter molecules decrease G protein activation. The specific response depends on which molecule is binding to the receptor.

One major group of bitter-tasting molecules are alkaloids. Alkaloids are nitrogen containing molecules that are commonly found in bitter-tasting plant products, such as coffee, hops (in beer), tannins (in wine), tea, and aspirin. By containing toxic alkaloids, the plant is less susceptible to microbe infection and less attractive to herbivores.

Therefore, the function of bitter taste may primarily be related to stimulating the gag reflex to avoid ingesting poisons. Because of this, many bitter foods that are normally ingested are often combined with a sweet component to make them more palatable (cream and sugar in coffee, for example). The highest concentration of bitter receptors appear to be in the posterior tongue, where a gag reflex could still spit out poisonous food.

The taste known as umami is often referred to as the savory taste. Like sweet and bitter, it is based on the activation of G protein–coupled receptors by a specific molecule. The molecule that activates this receptor is the amino acid L-glutamate. Therefore, the umami flavor is often perceived while eating protein-rich foods. Not surprisingly, dishes that contain meat are often described as savory.

Once the gustatory cells are activated by the taste molecules, they release neurotransmitters onto the dendrites of sensory neurons. These neurons are part of the facial and glossopharyngeal cranial nerves, as well as a component within the vagus nerve dedicated to the gag reflex. The facial nerve connects to taste buds in the anterior third of the tongue. The glossopharyngeal nerve connects to taste buds in the posterior two thirds of the tongue. The vagus nerve connects to taste buds in the extreme posterior of the tongue, verging on the pharynx, which are more sensitive to noxious stimuli such as bitterness.

The Tongue. The tongue is covered with small bumps, called papillae, which contain taste buds that are sensitive to chemicals in ingested food or drink. Different types of papillae are found in different regions of the tongue. The taste buds contain specialized gustatory receptor cells that respond to chemical stimuli dissolved in the saliva. These receptor cells activate sensory neurons that are part of the facial and glossopharyngeal nerves. LM × 1600. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)

Once the gustatory cells are activated by the taste molecules, they release neurotransmitters onto the dendrites of sensory neurons. These neurons are part of the facial and glossopharyngeal cranial nerves, as well as a component within the vagus nerve dedicated to the gag reflex. The facial nerve connects to taste buds in the anterior third of the tongue. The glossopharyngeal nerve connects to taste buds in the posterior two thirds of the tongue. The vagus nerve connects to taste buds in the extreme posterior of the tongue, verging on the pharynx, which are more sensitive to noxious stimuli such as bitterness.

Nerves of the Face and Oral Cavity

An iconic part of a doctor’s visit is the inspection of the oral cavity and pharynx, suggested by the directive to “open your mouth and say ‘ah.’” This is followed by inspection, with the aid of a tongue depressor, of the back of the mouth, or the opening of the oral cavity into the pharynx known as the fauces. Whereas this portion of a medical exam inspects for signs of infection, such as in tonsillitis, it is also the means to test the functions of the cranial nerves that are associated with the oral cavity.

The facial and glossopharyngeal nerves convey gustatory stimulation to the brain. Testing this is as simple as introducing salty, sour, bitter, or sweet stimuli to either side of the tongue. The patient should respond to the taste stimulus before retracting the tongue into the mouth. Stimuli applied to specific locations on the tongue will dissolve into the saliva and may stimulate taste buds connected to either the left or right of the nerves, masking any lateral deficits. Along with taste, the glossopharyngeal nerve relays general sensations from the pharyngeal walls. These sensations, along with certain taste stimuli, can stimulate the gag reflex. If the examiner moves the tongue depressor to contact the lateral wall of the fauces, this should elicit the gag reflex. Stimulation of either side of the fauces should elicit an equivalent response. The motor response, through contraction of the muscles of the pharynx, is mediated through the vagus nerve. Normally, the vagus nerve is considered autonomic in nature. The vagus nerve directly stimulates the contraction of skeletal muscles in the pharynx and larynx to contribute to the swallowing and speech functions. Further testing of vagus motor function has the patient repeating consonant sounds that require movement of the muscles around the fauces. The patient is asked to say “lah-kah-pah” or a similar set of alternating sounds while the examiner observes the movements of the soft palate and arches between the palate and tongue.

The facial and glossopharyngeal nerves are also responsible for the initiation of salivation. Neurons in the salivary nuclei of the medulla project through these two nerves as preganglionic fibers, and synapse in ganglia located in the head. The parasympathetic fibers of the facial nerve synapse in the pterygopalatine ganglion, which projects to the submandibular gland and sublingual gland. The parasympathetic fibers of the glossopharyngeal nerve synapse in the otic ganglion, which projects to the parotid gland. Salivation in response to food in the oral cavity is based on a visceral reflex arc within the facial or glossopharyngeal nerves. Other stimuli that stimulate salivation are coordinated through the hypothalamus, such as the smell and sight of food.

The hypoglossal nerve is the motor nerve that controls the muscles of the tongue, except for the palatoglossus muscle, which is controlled by the vagus nerve. There are two sets of muscles of the tongue. The extrinsic muscles of the tongue are connected to other structures, whereas the intrinsic muscles of the tongue are completely contained within the lingual tissues. While examining the oral cavity, movement of the tongue will indicate whether hypoglossal function is impaired. The test for hypoglossal function is the “stick out your tongue” part of the exam. The genioglossus muscle is responsible for protrusion of the tongue. If the hypoglossal nerves on both sides are working properly, then the tongue will stick straight out. If the nerve on one side has a deficit, the tongue will stick out to that side—pointing to the side with damage. Loss of function of the tongue can interfere with speech and swallowing. Additionally, because the location of the hypoglossal nerve and nucleus is near the cardiovascular center, inspiratory and expiratory areas for respiration, and the vagus nuclei that regulate digestive functions, a tongue that protrudes incorrectly can suggest damage in adjacent structures that have nothing to do with controlling the tongue.

Motor Nerves of the Neck

The accessory nerve, also referred to as the spinal accessory nerve, innervates the sternocleidomastoid and trapezius muscles (Figure 16.11). When both the sternocleidomastoids contract, the head flexes forward; individually, they cause rotation to the opposite side. The trapezius can act as an antagonist, causing extension and hyperextension of the neck. These two superficial muscles are important for changing the position of the head. Both muscles also receive input from cervical spinal nerves. Along with the spinal accessory nerve, these nerves contribute to elevating the scapula and clavicle through the trapezius, which is tested by asking the patient to shrug both shoulders, and watching for asymmetry. For the sternocleidomastoid, those spinal nerves are primarily sensory projections, whereas the trapezius also has lateral insertions to the clavicle and scapula, and receives motor input from the spinal cord. Calling the nerve the spinal accessory nerve suggests that it is aiding the spinal nerves. Though that is not precisely how the name originated, it does help make the association between the function of this nerve in controlling these muscles and the role these muscles play in movements of the trunk or shoulders.

Figure 16.11 Muscles Controlled by the Accessory Nerve The accessory nerve innervates the sternocleidomastoid and trapezius muscles, both of which attach to the head and to the trunk and shoulders. They can act as antagonists in head flexion and extension, and as synergists in lateral flexion toward the shoulder.

To test these muscles, the patient is asked to flex and extend the neck or shrug the shoulders against resistance, testing the strength of the muscles. Lateral flexion of the neck toward the shoulder tests both at the same time. Any difference on one side versus the other would suggest damage on the weaker side. These strength tests are common for the skeletal muscles controlled by spinal nerves and are a significant component of the motor exam. Deficits associated with the accessory nerve may have an effect on orienting the head, as described with the VOR.

The hypoglossal nerve is the motor nerve that controls the muscles of the tongue, except for the palatoglossus muscle, which is controlled by the vagus nerve. There are two sets of muscles of the tongue. The extrinsic muscles of the tongue are connected to other structures, whereas the intrinsic muscles of the tongue are completely contained within the lingual tissues. While examining the oral cavity, movement of the tongue will indicate whether hypoglossal function is impaired. The test for hypoglossal function is the “stick out your tongue” part of the exam. The genioglossus muscle is responsible for protrusion of the tongue. If the hypoglossal nerves on both sides are working properly, then the tongue will stick straight out. If the nerve on one side has a deficit, the tongue will stick out to that side—pointing to the side with damage. Loss of function of the tongue can interfere with speech and swallowing. Additionally, because the location of the hypoglossal nerve and nucleus is near the cardiovascular center, inspiratory and expiratory areas for respiration, and the vagus nuclei that regulate digestive functions, a tongue that protrudes incorrectly can suggest damage in adjacent structures that have nothing to do with controlling the tongue.